The role of the annexin A2 heterotetramer in vascular fibrinolysis

Blood

Volumn 118, Issue 18, 2011, Pages 4789-4797

  Madureira, Patricia A ^a   Surette, Alexi P ^a   Phipps, Kyle D ^a   Taboski, Michael A S ^a   Miller, Victoria A ^a   Waisman, David M ^a,b  

^a DALHOUSIE UNIVERSITY   (Canada)

^b DALHOUSIE UNIVERSITY   (Canada)

Author keywords

[No Author keywords available]

Indexed keywords

CELL SURFACE RECEPTOR; LIPOCORTIN 2; PLASMIN; PLASMINOGEN; PROTEIN SUBUNIT; PROTEINASE; RECOMBINANT PROTEIN; TISSUE PLASMINOGEN ACTIVATOR; UROKINASE;

CELL MEMBRANE; CHEMOTAXIS; ELECTRON TOMOGRAPHY; FIBRINOLYSIS; FIBROSARCOMA; HUMAN; NONHUMAN; PLASMINOGEN ACTIVATION; PRIORITY JOURNAL; PROMYELOCYTIC LEUKEMIA; PROTEIN BINDING; PROTEIN BLOOD LEVEL; PROTEIN CONFORMATION; PROTEIN DEGRADATION; PROTEIN EXPRESSION; PROTEIN INTERACTION; PROTEIN STRUCTURE; REVIEW; VASCULAR ENDOTHELIUM; WESTERN BLOTTING;

EID: 80855133519     PISSN: 00064971     EISSN: 15280020     Source Type: Journal    
DOI: 10.1182/blood-2011-06-334672     Document Type: Review

References (113)

1. Augustin HG, Kozian DH, Johnson RC. Differentiation of endothelial cells: analysis of the constitutive and activated endothelial cell phenotypes. Bioessays. 1994;16(12):901-906. (Pubitemid 24974469)
2. Cines DB, Pollak ES, Buck CA, et al. Endothelial cells in physiology and in the pathophysiology of vascular disorders. Blood. 1998;91(10):3527-3561. (Pubitemid 28225719)
3. Vallet B. Bench-to-bedside review. Endothelial cell dysfunction in severe sepsis: a role in organ dysfunction? Crit Care. 2003;7(2):130-138. (Pubitemid 36432671)
4. Tanaka KA, Key NS, Levy JH. Blood coagulation: hemostasis and thrombin regulation. Anesth Analg. 2009;108(5):1433-1446.
5. Ait-Oufella H, Maury E, Lehoux S, Guidet B, Offenstadt G. The endothelium: physiological functions and role in microcirculatory failure during severe sepsis. Intensive Care Med. 2010;36(8):1286-1298.
6. Henschen A, Lottspeich F, Kehl M, Southan C. Covalent structure of fibrinogen. Ann N Y Acad Sci. 1983;408(1):28-43. (Pubitemid 13072024)
7. Mosesson MW. Fibrinogen and fibrin structure and functions. J Thromb Haemost. 2005;3(8):1894-1904. (Pubitemid 41716577)
8. Cilia La Corte AL, Philippou H, Ariëns RAS. Role of fibrin structure in thrombosis and vascular disease. Adv Protein Chem Struct Biol. 2011;83:75-127.
9. Roberts WW, Kramer O, Rosser RW, Nestler FH, Ferry JD. Rheology of fibrin clots. I. Dynamic viscoelastic properties and fluid permeation. Biophys Chem. 1974;1(3):152-160.
10. Lijnen HR, Collen D. Mechanisms of physiological fibrinolysis [review]. Baillieres Clin Haematol. 1995;8(2):277-290.
11. Vassalli JD, Sappino AP, Belin D. The plasminogen activator/plasmin system. J Clin Invest. 1991;88(4):1067-1072.
12. Plow EF, Felez J, Miles LA. Cellular regulation of fibrinolysis [review]. Thromb Haemost. 1991;66(1):32-36.
13. Gyetko MR, Shollenberger SB, Sitrin RG. Urokinase expression in mononuclear phagocytes: cytokine-specific modulation by interferon-gamma and tumor necrosis factor-alpha. J Leukoc Biol. 1992;51(3):256-263.
14. Gyetko MR, Wilkinson CC, Sitrin RG. Monocyte urokinase expression: modulation by interleukins. J Leukoc Biol. 1993;53(5):598-601. (Pubitemid 23190049)
15. Fay WP, Garg N, Sunkar M. Vascular functions of the plasminogen activation system. Arterioscler Thromb Vasc Biol. 2007;27(6):1231-1237. (Pubitemid 46809410)
16. Plow EF, Hoover-Plow J. The functions of plasminogen in cardiovascular disease. Trends Cardiovasc Med. 2004;14(5):180-186. (Pubitemid 38942825)
17. Ellis V. Plasminogen activation at the cell surface. Curr Top Dev Biol. 2003;54:263-312. (Pubitemid 137639472)
18. Miles LA, Castellino FJ, Gong Y. Critical role for conversion of Glu-plasminogen to Lys-plasminogen for optimal stimulation of plasminogen activation on cell surfaces. Trends Cardiovasc Med. 2003;13(1):21-30. (Pubitemid 36120546)
19. Miles LA, Hawley SB, Baik N, et al. Plasminogen receptors: the sine qua non of cell surface plasminogen activation. Front Biosci. 2005;10:1754-1762. (Pubitemid 40529035)
20. Redlitz A, Plow EF. Receptors for plasminogen and t-PA: an update. Baillieres Clin Haematol. 1995;8(2):313-327.
21. Fogg DK, Choi KS, Waisman DM. Plasminogen receptors. In: Plasminogen: Structure, Activation and Regulation. New York, NY: Kluwer Academic/Plenum; 2003:81-101.
22. Plow EF, Freaney DE, Plescia J, Miles LA. The plasminogen system and cell surfaces: evidence for plasminogen and urokinase receptors on the same cell type. J Cell Biol. 1986;103(6):2411-2420. (Pubitemid 17216665)
23. Ellis V, Behrendt N, Dano K. Plasminogen activation by receptor-bound urokinase: a kinetic study with both cell-associated and isolated receptor. J Biol Chem. 1991;266(19):12752-12758. (Pubitemid 21907159)
24. Gonzalez-Gronow M, Stack S, Pizzo SV. Plasmin binding to the plasminogen receptor enhances catalytic efficiency and activates the receptor for subsequent ligand binding. Arch Biochem Biophys. 1991;286(2):625-628.
25. Hall SW, Humphries JE, Gonias SL. Inhibition of cell surface receptor-bound plasmin by alpha 2-antiplasmin and alpha 2-macroglobulin. J Biol Chem. 1991;266(19):12329-12336. (Pubitemid 21907096)
26. Plow EF, Herren T, Redlitz A, Miles LA, Hoover-Plow JL. The cell biology of the plasminogen system [review]. FASEB J. 1995;9(10):939-945.
27. Felez J. Plasminogen binding to cell surfaces. Fibrinolysis Proteol. 1998;12:183-189. (Pubitemid 28539225)
28. Herren T, Swaisgood C, Plow EF. Regulation of plasminogen receptors. Front Biosci. 2003;8:D1-D8. (Pubitemid 39089046)
29. Kwon M, MacLeod TJ, Zhang Y, Waisman DM. S100A10, annexin A2, and annexin A2 heterotetramer as candidate plasminogen receptors. Front Biosci. 2005;10:300-325. (Pubitemid 40543035)
30. Felez J, Chanquia CJ, Levin EG, Miles LA, Plow EF. Binding of tissue plasminogen activator to human monocytes and monocytoid cells. Blood. 1991;78(9):2318-2327.
31. Felez J, Chanquia CJ, Fabregas P, Plow EF, Miles LA. Competition between plasminogen and tissue plasminogen activator for cellular binding sites. Blood. 1993;82(8):2433-2441. (Pubitemid 23304267)
32. Cesarman GM, Guevara CA, Hajjar KA. An endothelial cell receptor for plasminogen/tissue plasminogen activator (t-PA): II. Annexin II-mediated enhancement of t-PA- dependent plasminogen activation. J Biol Chem. 1994;269(33):21198-21203.
33. Hajjar KA, Jacovina AT, Chacko J. An endothelial cell receptor for plasminogen/tissue plasminogen activator: I. Identity with annexin II. J Biol Chem. 1994;269(33):21191-21197.
34. Hajjar KA, Mauri L, Jacovina AT, et al. Tissue plasminogen activator binding to the annexin II tail domain: direct modulation by homocysteine. J Biol Chem. 1998;273(16):9987-9993. (Pubitemid 28183100)
35. Kassam G, Choi KS, Ghuman J, et al. The role of annexin II tetramer in the activation of plasminogen. J Biol Chem. 1998;273(8):4790-4799. (Pubitemid 28103230)
36. Kassam G, Le BH, Choi KS, et al. The p11 subunit of the annexin II tetramer plays a key role in the stimulation of t-PA-dependent plasminogen activation. Biochemistry. 1998;37(48):16958-16966. (Pubitemid 28566773)
37. Choi KS, Fogg DK, Yoon CS, Waisman DM. P11 regulates extracellular plasmin production and invasiveness of HT1080 fibrosarcoma cells. FASEB J. 2003;17:235-246. (Pubitemid 36164393)
38. Zhang L, Fogg DK, Waisman DM. RNA interference-mediated silencing of the S100A10 gene attenuates plasmin generation and invasiveness of Colo 222 colorectal cancer cells. J Biol Chem. 2004;279(3):2053-2062. (Pubitemid 38084477)
39. O'Connell PA, Surette AP, Liwski RS, Svenningsson P, Waisman DM. S100A10 regulates plasminogen-dependent macrophage invasion. Blood. 2010;116(7):1136- 1146.
40. O'Connell PA, Madureira PA, Berman JN, Liwski RS, Waisman DM. Regulation of S100A10 by the PML-RAR-alpha oncoprotein. Blood. 2011;117(15):4095-4105.
41. Surette AP, Madureira PA, Phipps KD, et al. Regulation of fibrinolysis by S100A10 in vivo. Blood. 2011;118(11):3172-3181.
42. Fogg DK, Bridges DE, Cheung KT, et al. The p11 subunit of annexin II heterotetramer is regulated by basic carboxypeptidase. Biochemistry. 2002;41(15):4953-4961. (Pubitemid 34298663)
43. MacLeod TJ, Kwon M, Filipenko NR, Waisman DM. Phospholipid-associated annexin A2-S100A10 heterotetramer and its subunits: characterization of the interaction with tissue plasminogen activator, plasminogen, and plasmin. J Biol Chem. 2003;278(28):25577-25584. (Pubitemid 36835310)
44. Ling Q, Jacovina AT, Deora A, et al. Annexin II regulates fibrin homeostasis and neoangiogenesis in vivo. J Clin Invest. 2004;113(1):38-48. (Pubitemid 38541630)
45. Puisieux A, Ji J, Ozturk M. Annexin II up-regulates cellular levels of p11 protein by a post-translational mechanisms. Biochem J. 1996;313(1):51-55. (Pubitemid 26013424)
46. Zhao WQ, Chen GH, Chen H, et al. Secretion of Annexin II via activation of insulin receptor and insulin-like growth factor receptor. J Biol Chem. 2003;278(6):4205-4215. (Pubitemid 36801158)
47. Deora AB, Kreitzer G, Jacovina AT, Hajjar KA. An annexin 2 phosphorylation switch mediates p11-dependent translocation of annexin 2 to the cell surface. J Biol Chem. 2004;279(42):43411-43418. (Pubitemid 39390640)
48. Valapala M, Vishwanatha JK. Lipid raft endocytosis and exosomal transport facilitate extracellular trafficking of annexin A2. http://www.ncbi.nlm.nih. gov/pubmed/21737841. Accessed July 29, 2011.
49. Zheng L, Foley K, Huang L, et al. Tyrosine 23 phosphorylation-dependent cell-surface localization of annexin A2 is required for invasion and metastases of pancreatic cancer. PLoS One. 2011;6(4):e19390.
50. Filipenko NR, Waisman DM. Annexin II: analysis of a pleiotropic protein. In: Annexins: Biological Importance and Annexin-Related Pathologies. New York, NY: Landes Bioscience; 2003;127-156.
51. Gerke V, Moss SE. Annexins: from structure to function. Physiol Rev. 2002;82(2):331-371. (Pubitemid 34654456)
52. .Waisman DM. Annexin II tetramer: structure and function. Mol Cell Biochem. 1995;149:301-322.
53. Lambert O, Gerke V, Bader MF, Porte F, Brisson A. Structural analysis of junctions formed between lipid membranes and several annexins by cryoelectron microscopy. J Mol Biol. 1997;272(1):42-55. (Pubitemid 27395536)
54. Réty S, Sopkova J, Renouard M, et al. The crystal structure of a complex of p11 with the annexin II N-terminal peptide. Nat Struct Biol. 1999;6(1):89-95. (Pubitemid 29232429)
55. Becker T, Weber K, Johnsson N. Protein-protein recognition via short amphiphilic helices: a mutational analysis of the binding site of annexin II for p11. EMBO J. 1990;9(13):4207-4213. (Pubitemid 120025975)
56. Kube E, Becker T, Weber K, Gerke V. Protein-protein interaction studied by site-directed mutagenesis: characterization of the annexin II-binding site on p11, a member of the S100 protein family. J Biol Chem. 1992;267(20):14175- 14182.
57. Rescher U, Gerke V. S100A10/p11: family, friends and functions. Pflugers Arch. 2007;455(4):575-582. (Pubitemid 350160469)
58. Donato R. S100: a multigenic family of calcium-modulated proteins of the EF-hand type with intracellular and extracellular functional roles. Int J Biochem Cell Biol. 2001;33(7):637-668. (Pubitemid 32520962)
59. Semov A, Moreno MJ, Onichtchenko A, et al. Metastasis-associated protein S100A4 induces angiogenesis through interaction with Annexin II and accelerated plasmin formation. J Biol Chem. 2005;280(21):20833-20841. (Pubitemid 40776788)
60. Zeng FY, Gerke V, Gabius HJ. Identification of annexin II, annexin VI and glyceraldehyde-3-phosphate dehydrogenase as calcyclin-binding proteins in bovine heart. Int J Biochem. 1993;25(7):1019-1027. (Pubitemid 23186507)
61. Johnsson N, Vandekerckhove J, Van Damme J, Weber K. Binding sites for calcium, lipid and p11 on p36, the substrate of retroviral tyrosine-specific protein kinases. FEBS Lett. 1986;198(2):361-364.
62. Mailliard WS, Haigler HT, Schlaepfer DD. Calcium-dependent binding of S100C to the N-terminal domain of annexin I. J Biol Chem. 1996;271(2):719-725. (Pubitemid 26034919)
63. Garbuglia M, Verzini M, Donato R. Annexin VI binds S100A1 and S100B and blocks the ability of S100A1 and S100B to inhibit desmin and GFAP assemblies into intermediate filaments. Cell Calcium. 1998;24(3):177-191. (Pubitemid 28552188)
64. Watanabe M, Ando Y, Tokumitsu H, Hidaka H. Binding site of annexin XI on the calcyclin molecule. Biochem Biophys Res Commun. 1993;196(3):1376-1382. (Pubitemid 23347602)
65. Gerke V, Weber K. Identity of p36K phosphorylated upon Rous sarcoma virus transformation with a protein purified from brush borders: calcium-dependent binding to non-erythroid spectrin and F-actin. EMBO J. 1984;3:227-233.
66. Zokas L, Glenney JR. The calpactin light chain is tightly linked to the cytoskeletal form of calpactin I: studies using monoclonal antibodies to calpactin subunits. J Cell Biol. 1987;105:2111-2121. (Pubitemid 18019062)
67. He K-L, Deora AB, Xiong H, et al. Endothelial cell annexin A2 regulates polyubiquitination and degradation of its binding partner S100A10/p11. J Biol Chem. 2008;283(28):19192-19200.
68. Gladwin MT, Yao XL, Cowan M, et al. Retinoic acid reduces p11 protein levels in bronchial epithelial cells by a posttranslational mechanism. Am J Physiol Lung Cell Mol Physiol. 2000;279(6):L1103-L1109. (Pubitemid 32009496)
69. Gerke V, Creutz CE, Moss SE. Annexins: linking Ca2+ signalling to membrane dynamics. Nat Rev Mol Cell Biol. 2005;6(6):449-461. (Pubitemid 40780557)
70. Moss SE, Morgan RO. The annexins. Genome Biol. 2004;5(4):219.
71. Rescher U, Gerke V. Annexins: unique membrane binding proteins with diverse functions. J Cell Sci. 2004;117(13):2631-2639. (Pubitemid 38997247)
72. Rintala-Dempsey AC, Rezvanpour A, Shaw GS. S100-annexin complexes: structural insights. FEBS J. 2008;275(20):4956-4966.
73. Burger A, Berendes R, Liemann S, et al. The crystal structure and ion channel activity of human Annexin II, a peripheral membrane protein. J Mol Biol. 1996;257:839-847. (Pubitemid 26117072)
74. Arrigo AP, Darlix JL, Spahr PF. A cellular protein phosphorylated by the avian sarcoma virus transforming gene product is associated with ribonucleoprotein particles. EMBO J. 1983;2(3):309-315.
75. Tressler RJ, Updyke TV, Yeatman T, Nicolson GL. Extracellular annexin II is associated with divalent cation-dependent tumor cell-endothelial cell adhesion of metastatic RAW117 large-cell lymphoma cells. J Cell Biochem. 1993;53:265-276. (Pubitemid 23323111)
76. Courtneidge S, Ralston R, Alitalo K, Bishop JM. Subcellular location of an abundant substrate (p36) for tyrosine-specific protein kinases. Mol Cell Biol. 1983;3(3):340-350. (Pubitemid 13165199)
77. Vedeler A, Hollas H. Annexin II is associated with mRNAs which may constitute a distinct subpopulation. Biochem J. 2000;348(3):565-572.
78. MacLeod TJ, Filipenko NR, Waisman DM. Characterization of the RNA-binding properties of annexin A2 heterotetramer. Annexins. 2004;1(3):191-199.
79. Venetsanakos E, Mirza A, Fanton C, et al. Induction of tubulogenesis in telomerase-immortalized human microvascular endothelial cells by glioblastoma cells. Exp Cell Res. 2002;273(1):21-33.
80. Hou Y, Yang L, Mou M, et al. Annexin A2 regulates the levels of plasmin, S100A10 and fascin in L5178Y Cells. Cancer Invest. 2008;26(8):809-815.
81. Zobiack N, Rescher U, Ludwig C, Zeuschner D, Gerke V. The annexin 2/S100A10 complex controls the distribution of transferrin receptor-containing recycling endosomes. Mol Biol Cell. 2003;14(12):4896-4908. (Pubitemid 37484805)
82. Mai J, Waisman DM, Sloane BF. Cell surface complex of cathepsin B/annexin II tetramer in malignant progression. Biochim Biophys Acta. 2000;1477(1):215-230. (Pubitemid 30119902)
83. Swisher JFA, Burton N, Bacot SM, Vogel SN, Feldman GM. Annexin A2 tetramer activates human and murine macrophages through TLR4. Blood. 2010;115(3):549-558.
84. Kapoor M, Zhou Q, Otero F, et al. Evidence for Annexin II-S100A10 complex and plasmin in mobilization of cytokine activity of human TrpRS. J Biol Chem. 2008;283(4):2070-2077.
85. Laumonnier Y, Syrovets T, Burysek L, Simmet T. Identification of the annexin A2 heterotetramer as a receptor for the plasmin-induced signaling in human peripheral monocytes. Blood. 2006;107(8):3342-3349.
86. .Waisman DM. Annexin A2 may not play a role as a plasminogen receptor. Br J Haematol. 2005;131(4):553-554.
87. Kang HM, Kassam G, Jarvis SE, Fitzpatrick SL, Waisman DM. Characterization of human recombinant annexin II tetramer purified from bacteria: role of N-terminal acetylation. Biochemistry. 1997;36:2041-2050. (Pubitemid 27104694)
88. Galántai R, Módos K, Fidy J, Kolev K, Machovich R. Structural basis of the cofactor function of denatured albumin in plasminogen activation by tissue-type plasminogen activator. Biochem Biophys Res Commun. 2006;341(3):736-741. (Pubitemid 43177551)
89. Machovich R, Owen WG. Denatured proteins as cofactors for plasminogen activation. Arch Biochem Biophys. 1997;344:343-349. (Pubitemid 27344071)
90. Radcliffe R, Heinze T. Stimulation of tissue plasminogen activator by denatured proteins and fibrin clots: a possible additional role for plasminogen activator? Arch Biochem Biophys. 1981;211(2):750-761. (Pubitemid 12157849)
91. Jost M, Weber K, Gerke V. Annexin II contains two types of Ca(2+)-binding sites. Biochem J. 1994;298(3):553-559. (Pubitemid 24085176)
92. Das R, Burke T, Plow EF. Histone H2B as a functionally important plasminogen receptor on macrophages. Blood. 2007;110(10):3763-3772. (Pubitemid 350159647)
93. Hajjar KA, Hamel NM. Identification and characterization of human endothelial cell membrane binding sites for tissue plasminogen activator and urokinase. J Biol Chem. 1990;265(5):2908-2916.
94. Felez J, Miles LA, Fabregas P, et al. Characterization of cellular binding sites and interactive regions within reactants required for enhancement of plasminogen activation by tPA on the surface of leukocytic cells. Thromb Haemost. 1996;76(4):577-584. (Pubitemid 26372014)
95. Roda O, Valero ML, Peiró S, et al. New insights into the tPA-annexin A2 interaction: is annexin A2 CYS8 the sole requirement for this association? J Biol Chem. 2003;278(8):5702-5709. (Pubitemid 36800814)
96. Stepchenko AG, Luchina NN, Pankratova EV. Cysteine 50 of the POUH domain determines the range of targets recognized by POU proteins. Nucleic Acids Res. 1997;25(14):2847-2853. (Pubitemid 27299801)
97. Helmstaedt K, Krappmann S, Braus GH. Allosteric regulation of catalytic activity: Escherichia coli aspartate transcarbamoylase versus yeast chorismate mutase. Microbiol Mol Biol Rev. 2001;65(3):404-421. (Pubitemid 32881713)
98. Stephens RW, Pollanen J, Tapiovaara H, et al. Activation of pro-urokinase and plasminogen on human sarcoma cells: a proteolytic system with surface-bound reactants. J Cell Biol. 1989;108(5):1987-1995. (Pubitemid 19130529)
99. Mai J, Finley RL, Waisman DM, Sloane BF. Human procathepsin B interacts with the annexin II tetramer on the surface of tumor cells. J Biol Chem. 2000;275(17):12806-12812. (Pubitemid 30241435)
100. Kwaan HC. The plasminogen-plasmin system in malignancy [review]. Cancer Metastasis Rev. 1992;11(3):291-311.
101. Del Rosso M, Fibbi G, Pucci M, et al. Multiple pathways of cell invasion are regulated by multiple families of serine proteases. Clin Exp Metastasis. 2002;19(3):193-207. (Pubitemid 34553521)
102. Smith HW, Marshall CJ. Regulation of cell signalling by uPAR. Nat Rev Mol Cell Biol. 2010;11(1):23-36.
103. Blasi F, Sidenius N. The urokinase receptor: focused cell surface proteolysis, cell adhesion and signaling. FEBS Lett. 2010;584(9):1923-1930.
104. Gong Y, Hart E, Shchurin A, Hoover-Plow J. Inflammatory macrophage migration requires MMP-9 activation by plasminogen in mice. J Clin Invest. 2008;118(9):3012-3024.
105. Ploplis VA, French EL, Carmeliet P, Collen D, Plow EF. Plasminogen deficiency differentially affects recruitment of inflammatory cell populations in mice. Blood. 1998;91(6):2005-2009. (Pubitemid 28185501)
106. Tallman MS, Abutalib SA, Altman JK. The double hazard of thrombophilia and bleeding in acute promyelocytic leukemia. Semin Thromb Hemost. 2007;33(4):330-338. (Pubitemid 46878105)
107. Avvisati G, Lo Coco F, Mandelli F. Acute promyelocytic leukemia: clinical and morphologic features and prognostic factors. Semin Hematol. 2001;38(1):4-12. (Pubitemid 32107698)
108. Arbuthnot C, Wilde JT. Haemostatic problems in acute promyelocytic leukaemia. Blood Rev. 2006;20(6):289-297. (Pubitemid 44692001)
109. Menell JS, Cesarman GM, Jacovina AT, et al. Annexin II and bleeding in acute promyelocytic leukemia. N Engl J Med. 1999;340(13):994-1004. (Pubitemid 29157631)
110. Miles LA, Ellis V. Alpha-enolase comes muscling in on plasminogen activation. Thromb Haemost. 2003;90(4):564-566. (Pubitemid 37287621)
111. Miles LA, Dahlberg CM, Plescia J, et al. Role of cell-surface lysines in plasminogen binding to cells: identification of alpha-enolase as a candidate plasminogen receptor. Biochemistry. 1991;30(6):1682-1691.
112. Hawley SB, Tamura T, Miles LA. Purification, cloning, and characterization of a profibrinolytic plasminogen-binding protein, TIP49a. J Biol Chem. 2001;276(1):179-186. (Pubitemid 32050305)
113. Andronicos NM, Chen EI, Baik N, et al. Proteomics-based discovery of a novel, structurally unique, and developmentally regulated plasminogen receptor, Plg-RKT, a major regulator of cell surface plasminogen activation. Blood. 2010;115(7):1319-1330.